Ecosystem growth and development.

One of the most important features of biosystems is how they are able to maintain local order (low entropy) within their system boundaries. At the ecosystem scale, this organization can be observed in the thermodynamic parameters that describe it, such that these parameters can be used to track ecosystem growth and development during succession. Thermodynamically, ecosystem growth is the increase of energy throughflow and stored biomass, and ecosystem development is the internal reorganization of these energy mass stores, which affect transfers, transformations, and time lags within the system. Several proposed hypotheses describe thermodynamically the orientation or natural tendency that ecosystems follow during succession, and here, we consider five: minimize specific entropy production, maximize dissipation, maximize exergy storage (includes biomass and information), maximize energy throughflow, and maximize retention time. These thermodynamic orientors were previously all shown to occur to some degree during succession, and here we present a refinement by observing them during different stages of succession. We view ecosystem succession as a series of four growth and development stages: boundary, structural, network, and informational. We demonstrate how each of these ecological thermodynamic orientors behaves during the different growth and development stages, and show that while all apply during some stages only maximizing energy throughflow and maximizing exergy storage are applicable during all four stages. Therefore, we conclude that the movement away from thermodynamic equilibrium, and the subsequent increase in organization during ecosystem growth and development, is a result of system components and configurations that maximize the flux of useful energy and the amount of stored exergy. Empirical data and theoretical models support these conclusions.